Hind limb stepping over obstacles in the horse guided by place-object memory

An animal that has stepped over an obstacle with its forelimbs uses a memory of the obstacle to guide the hind limbs so that they also clear the obstacle, even in situations in which long pauses are introduced between forelimb and hind limb stepping. To further clarify the features of hind limb obstacle clearance memory, the present study examined hind limb obstacle clearance in the horse. A rider guided horses over obstacles and paused the horse over obstacles in tests that examined the relationship between forelimb and hind limb stepping, with the following results. First, the horses displayed memory for an obstacle as measured by hind limb lifting over the obstacle for durations lasting as long as 15 min. The response was not dependent upon ongoing visualization of the obstacle, as limb lifting was unaffected by visual occlusion with blinders, a blindfold, or by removing the obstacle during the pause. Second, previous experience of stepping over an obstacle led to pause-related hind limb lifting at the object's previous location even on trials for which there was no obstacle and so no preceding forelimb lifting. Third, whereas a horse would lift its hind limbs to clear two successively presented obstacles, replacing an obstacle before the horse after the forelimbs had cleared the obstacle prevented subsequent hind limb lifting at the obstacle's previous location. Taken together the results show that hind limb obstacle clearance is guided by a place-object memory. The results are discussed in relation to the differential sensory and memonic control of forelimb and hind limb stepping with the suggestion that place-object memory can guide hind stepping as well as overshadow working memory from front leg stepping.